Invertebrate

The common fruit fly, Drosophila melanogaster, has been used extensively for research

Invertebrates are animals that neither possess nor develop a vertebral column, derived from the notochord. This includes all animals apart from the subphylum Vertebrata. Familiar examples of invertebrates include insects; crabs, lobsters and their kin; snails, clams, octopuses and their kin; starfish, sea-urchins and their kin; and worms.

The majority of animal species are invertebrates; one estimate puts the figure at 97%. Many invertebrate taxa have a greater number and variety of species than the entire subphylum of Vertebrata.

Some of the so-called invertebrates, such as the Chaetognatha, Hemichordata, Tunicata and Cephalochordata are more closely related to the vertebrates than to other invertebrates. This makes the term "invertebrate" paraphyletic and hence almost meaningless for taxonomic purposes.

Etymology

The word "invertebrate" comes from the form of the Latin word vertebra, which means a joint in general, and sometimes specifically a joint from the spinal column of a vertebrate. In turn the jointed aspect of vertebra derived from the concept of turning, expressed in the root verto or vorto, to turn. Coupled with the prefix in-, meaning "not" or "without".

Taxonomic significance

The term invertebrates is not always precise among non-biologists since it does not accurately describe a taxon in the same way that Arthropoda, Vertebrata or Manidae do. Each of these terms describes a valid taxon, phylum, subphylum or family. "Invertebrata" is a term of convenience, not a taxon; it has very little circumscriptional significance except within the Chordata. The Vertebrata as a subphylum comprises such a small proportion of the Metazoa that to speak of the kingdom Animalia in terms of "Vertebrata" and "Invertebrata" has limited practicality. In the more formal taxonomy of Animalia other attributes that logically should precede the presence or absence of the vertebral column in constructing a cladogram, for example, the presence of a notochord. That would at least circumscribe the Chordata. However, even the notochord would be a less fundamental criterion than aspects of embryological development and symmetry or perhaps bauplan.

Invertebrates don't have a skeleton of bone, either internal or external. They include hugely varied body plans. Many have fluid-filled, hydrostatic skeletons, like jellyfish or worms. Others have hard exoskeletons, outer shells like those of insects and crustaceans. The most familiar invertebrates include the Protozoa, Porifera, Coelenterata, Platyhelminthes, Nematoda, Annelida, Echinodermata, Mollusca and Arthropoda. Arthropoda include insects, crustaceans and arachnids.

Number of extant species

By far the largest number of described invertebrate species are insects. The following table lists the number of described extant species for each major invertebrate group as estimated in the IUCN Red List of Threatened Species, 2014.3.

Invertebrate group	Image	Estimated number of described species
Insects		1,000,000
Molluscs		85,000
Crustaceans		47,000
Corals		2,000
Arachnids		102,248
Velvet worms		165
Horseshoe crabs		4
Others		68,658
	Totals:	1,305,075

The IUCN estimates that 66,178 extant vertebrate species have been described, which means that over 95% of the described animal species in the world are invertebrates.

Characteristics

The trait that is common to all invertebrates is the absence of a vertebral column: this creates a distinction between invertebrates and vertebrates. The distinction is one of convenience only; it is not based on any clear biologically homologous trait, any more than the common trait of having wings functionally unites insects, bats, and birds, or than not having wings unites tortoises, snails and sponges. Being animals, invertebrates are heterotrophs, and require sustenance in the form of the consumption of other organisms. With a few exceptions, such as the Porifera, invertebrates generally have bodies composed of differentiated tissues. There is also typically a digestive chamber with one or two openings to the exterior.

Morphology

All gastropods are asymmetrical. This is easily seen in snails and sea snails, which have helical shells. At first glance slugs appear externally symmetrical, but their pneumostome (breathing hole) is always on the right side. The origin of asymmetry in gastropods is a subject of scientific debate. Other gastropods develop external asymmetry, such as Glaucus atlanticus that develops asymmetrical cerata as they mature.

Fiddler crabs and hermit crabs have one claw much larger than the other. If a male fiddler loses its large claw, it will grow another on the opposite side after moulting. Sessile animals such as sponges are asymmetrical.

Corals build colonies that are not symmetrical, but the individual polyps exhibit radial symmetry.

Alpheidae feature asymmetrical claws that lack pincers.

Certain polyopisthocotylean monogeneans are asymmetrical, as an adaptation to their attachment to the gill of their fish hosts. Certain parasitic copepods which live inside the gill chamber of their fish hosts are asymmetrical.

Nervous system

Neurons differ in invertebrates from mammalian cells. Invertebrates cells fire in response to similar stimuli as mammals, such as tissue trauma, high temperature, or changes in pH. The first invertebrate in which a neuron cell was identified was the medicinal leech, Hirudo medicinalis.

Learning and memory using nociceptors in the sea hare, Aplysia has been described. Mollusk neurons are able to detect increasing pressures and tissue trauma.

Neurons have been identified in a wide range of invertebrate species, including annelids, molluscs, nematodes and arthropods.

Respiratory system

Tracheal system of dissected cockroach. The largest tracheae run across the width of the body of the cockroach and are horizontal in this image. Scale bar, 2 mm.

The tracheal system branches into progressively smaller tubes, here supplying the crop of the cockroach. Scale bar, 2.0 mm.

One type of invertebrate respiriatory system is the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues. The distribution of spiracles can vary greatly among the many orders of insects, but in general each segment of the body can have only one pair of spiracles, each of which connects to an atrium and has a relatively large tracheal tube behind it. The tracheae are invaginations of the cuticular exoskeleton that branch (anastomose) throughout the body with diameters from only a few micrometres up to 0.8 mm. The smallest tubes, tracheoles, penetrate cells and serve as sites of diffusion for water, oxygen, and carbon dioxide. Gas may be conducted through the respiratory system by means of active ventilation or passive diffusion. Unlike vertebrates, insects do not generally carry oxygen in their haemolymph.

A tracheal tube may contain ridge-like circumferential rings of taenidia in various geometries such as loops or helices. In the head, thorax, or abdomen, tracheae may also be connected to air sacs. Many insects, such as grasshoppers and bees, which actively pump the air sacs in their abdomen, are able to control the flow of air through their body. In some aquatic insects, the tracheae exchange gas through the body wall directly, in the form of a gill, or function essentially as normal, via a plastron. Note that despite being internal, the tracheae of arthropods are shed during moulting (ecdysis).

Reproduction

Like vertebrates, most invertebrates reproduce at least partly through sexual reproduction. They produce specialized reproductive cells that undergo meiosis to produce smaller, motile spermatozoa or larger, non-motile ova. These fuse to form zygotes, which develop into new individuals. Others are capable of asexual reproduction, or sometimes, both methods of reproduction.

Social interaction

Social behavior is widespread in invertebrates, including cockroaches, termites, aphids, thrips, ants, bees, Passalidae, Acari, spiders, and more. Social interaction is particularly salient in eusocial species but applies to other invertebrates as well.

Insects recognize information transmitted by other insects.

Phyla

The fossil coral Cladocora from the Pliocene of Cyprus

The term invertebrates covers several phyla. One of these are the sponges (Porifera). They were long thought to have diverged from other animals early. They lack the complex organization found in most other phyla. Their cells are differentiated, but in most cases not organized into distinct tissues. Sponges typically feed by drawing in water through pores. Some speculate that sponges are not so primitive, but may instead be secondarily simplified. The Ctenophora and the Cnidaria, which includes sea anemones, corals, and jellyfish, are radially symmetric and have digestive chambers with a single opening, which serves as both the mouth and the anus. Both have distinct tissues, but they are not organized into organs. There are only two main germ layers, the ectoderm and endoderm, with only scattered cells between them. As such, they are sometimes called diploblastic.

The Echinodermata are radially symmetric and exclusively marine, including starfish (Asteroidea), sea urchins, (Echinoidea), brittle stars (Ophiuroidea), sea cucumbers (Holothuroidea) and feather stars (Crinoidea).

The largest animal phylum is also included within invertebrates: the Arthropoda, including insects, spiders, crabs, and their kin. All these organisms have a body divided into repeating segments, typically with paired appendages. In addition, they possess a hardened exoskeleton that is periodically shed during growth. Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share these traits. The Nematoda or roundworms, are perhaps the second largest animal phylum, and are also invertebrates. Roundworms are typically microscopic, and occur in nearly every environment where there is water. A number are important parasites. Smaller phyla related to them are the Kinorhyncha, Priapulida, and Loricifera. These groups have a reduced coelom, called a pseudocoelom. Other invertebrates include the Nemertea or ribbon worms, and the Sipuncula.

Another phylum is Platyhelminthes, the flatworms. These were originally considered primitive, but it now appears they developed from more complex ancestors. Flatworms are acoelomates, lacking a body cavity, as are their closest relatives, the microscopic Gastrotricha. The Rotifera or rotifers, are common in aqueous environments. Invertebrates also include the Acanthocephala or spiny-headed worms, the Gnathostomulida, Micrognathozoa, and the Cycliophora.

Also included are two of the most successful animal phyla, the Mollusca and Annelida. The former, which is the second-largest animal phylum by number of described species, includes animals such as snails, clams, and squids, and the latter comprises the segmented worms, such as earthworms and leeches. These two groups have long been considered close relatives because of the common presence of trochophore larvae, but the annelids were considered closer to the arthropods because they are both segmented. Now, this is generally considered convergent evolution, owing to many morphological and genetic differences between the two phyla.

Among lesser phyla of invertebrates are the Hemichordata, or acorn worms, and the Chaetognatha, or arrow worms. Other phyla include Acoelomorpha, Brachiopoda, Bryozoa, Entoprocta, Phoronida, and Xenoturbellida.

Classification of invertebrates

Invertebrates can be classified into several main categories, some of which are taxonomically obsolescent or debatable, but still used as terms of convenience. Each however appears in its own article at the following links.

• Protozoa (like the worms, an arbitrary grouping of convenience; link to their article for details)
• Sponges (Porifera)
• Stinging jellyfish (Cnidaria)
• Comb jellies (Ctenophora)
• Flatworms (Platyhelminthes)
• Round- or threadworms (Nematoda)
• segmented worms (Annelida)
• Insects, spiders, crabs and their kin (Arthropoda)
• Cuttlefish, snails, mussels and their kin (Mollusca)
• Starfish, sea-cucumbers and their kin (Echinodermata)

History

The earliest animal fossils appear to be those of invertebrates. 665-million-year-old fossils in the Trezona Formation at Trezona Bore, West Central Flinders, South Australia have been interpreted as being early sponges. Some paleontologists suggest that animals appeared much earlier, possibly as early as 1 billion years ago. Trace fossils such as tracks and burrows found in the Tonian era indicate the presence of triploblastic worms, like metazoans, roughly as large (about 5 mm wide) and complex as earthworms.

Around 453 MYA, animals began diversifying, and many of the important groups of invertebrates diverged from one another. Fossils of invertebrates are found in various types of sediment from the Phanerozoic. Fossils of invertebrates are commonly used in stratigraphy.

Classification

Carl Linnaeus divided these animals into only two groups, the Insecta and the now-obsolete Vermes (worms). Jean-Baptiste Lamarck, who was appointed to the position of "Curator of Insecta and Vermes" at the Muséum National d'Histoire Naturelle in 1793, both coined the term "invertebrate" to describe such animals, and divided the original two groups into ten, by splitting Arachnida and Crustacea from the Linnean Insecta, and Mollusca, Annelida, Cirripedia, Radiata, Coelenterata and Infusoria from the Linnean Vermes. They are now classified into over 30 phyla, from simple organisms such as sea sponges and flatworms to complex animals such as arthropods and molluscs.

Significance of the group

Invertebrates are animals without a vertebral column. This has led to the conclusion that invertebrates are a group that deviates from the normal, vertebrates. This has been said to be because researchers in the past, such as Lamarck, viewed vertebrates as a "standard": in Lamarck's theory of evolution, he believed that characteristics acquired through the evolutionary process involved not only survival, but also progression toward a "higher form", to which humans and vertebrates were closer than invertebrates were. Although goal-directed evolution has been abandoned, the distinction of invertebrates and vertebrates persists to this day, even though the grouping has been noted to be "hardly natural or even very sharp." Another reason cited for this continued distinction is that Lamarck created a precedent through his classifications which is now difficult to escape from. It's also possible that some humans believe that, they themselves being vertebrates, the group deserves more attention than invertebrates. In any event, in the 1968 edition of Invertebrate Zoology, it is noted that "division of the Animal Kingdom into vertebrates and invertebrates is artificial and reflects human bias in favor of man's own relatives." The book also points out that the group lumps a vast number of species together, so that no one characteristic describes all invertebrates. In addition, some species included are only remotely related to one another, with some more related to vertebrates than other invertebrates (see Paraphyly).

In research

For many centuries, invertebrates have been neglected by biologists, in favor of big vertebrates and "useful" or charismatic species. Invertebrate biology did not get really coined until the XVIIIth century, with Linnaeus and Lamarck. During the XXth century, many discoveries made invertebrate zoology one of the major fields of natural sciences, with prominent discoveries for medicine, genetics, palaeontology, ecology, etc... Even law enforcement is now grateful to invertebrate science : arthropods, especially insects, are often used by forensic scientists, as some invertebrates are attracted to dead bodies.

Two of the most commonly studied model organisms nowadays are invertebrates: the fruit fly Drosophila melanogaster and the nematode Caenorhabditis elegans. They have long been the most intensively studied model organisms, and were among the first life-forms to be genetically sequenced. This was facilitated by the severely reduced state of their genomes, but many genes, introns, and linkages have been lost. Analysis of the starlet sea anemone genome has emphasised the importance of sponges, placozoans, and choanoflagellates, also being sequenced, in explaining the arrival of 1500 ancestral genes unique to animals. Invertebrates are also used by scientists in the field of aquatic biomonitoring to evaluate the effects of water pollution and climate change.

See also

• Invertebrate zoology
• Invertebrate paleozoology
• Marine invertebrates
• Pain in invertebrates

Further reading

• Hyman, L. H. 1940. The Invertebrates (6 volumes) New York : McGraw-Hill. A classic work.
• Anderson, D. T. (Ed.). (2001). Invertebrate zoology (2nd ed.). Oxford: Oxford University Press.
• Brusca, R. C., & Brusca, G. J. (2003). Invertebrates (2nd ed.). Sunderland, Mass. : Sinauer Associates.
• Miller, S.A., & Harley, J.P. (1996). Zoology (4th ed.). Boston: WCB/McGraw-Hill.
• Ruppert, E. E., Fox, R. S., & Barnes, R. D. (2004). Invertebrate zoology: a functional evolutionary approach. Belmont, CA: Thomas-Brooks/Cole.
• Adiyodi, K.G. & Adyiodi, R.G. (Eds) 1983- . Reproductive Biology of Invertebrates. Wiley, New York. (Many volumes.)
• Giese, A.G. & Pearse, J.S. (Eds) 1974- . Reproduction of Marine Invertebrates. Academic Press, New York. (Many volumes.)
• Advances in Invertebrate Reproduction. Elsevier Science, Amsterdam. (Five volumes.)

External links

• Buglife (UK)
• African Invertebrates